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Morfoanatomía Floral De Kallstroemia Maxima (Zygophyllaceae)

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Morfoanatomía Floral De Kallstroemia Maxima (Zygophyllaceae) Revista Mexicana de Biodiversidad Revista Mexicana de Biodiversidad 90 (2019): e902075 Anatomía Morfoanatomía floral de Kallstroemia maxima (Zygophyllaceae) Floral morphoanatomy of Kallstroemia maxima (Zygophyllaceae) Rosa María Fonseca a, *, Mercedes Eunice Castro-Laportte b y Estela Sandoval-Zapotitla c a Facultad de Ciencias, Universidad Nacional Autónoma de México, Circuito Exterior s/n, Ciudad Universitaria, 04510 Ciudad de México, México b Facultad de Agronomía, Universidad Central de Venezuela, Maracay 2101, Aragua, Venezuela c Jardín Botánico, Instituto de Biología, Universidad Nacional Autónoma de México, Tercer Circuito Exterior s/n, Ciudad Universitaria, 04510 Ciudad de México, México *Autor para correspondencia: [email protected] (R.M. Fonseca) Recibido: 9 enero 2016; aceptado: 30 julio 2017 Resumen El objetivo es contribuir al conocimiento de la anatomía floral de Kallstroemia maxima y compararlo con otros géneros relacionados dentro de la familia Zygophyllaceae. Se utilizaron botones florales de 3 plantas de K. maxima recolectados el día previo a la antesis; se procesaron preparaciones permanentes y se hicieron observaciones a partir de fotomicrografías de botones herborizados. Se proporcionan descripciones de algunas estructuras anatómicas no descritas con anterioridad, como son: nectarios opuestos a los sépalos, presencia de un haz vascular compartido por un pétalo y el estambre opuesto a éste, engrosamientos helicoidales en las células epidérmicas de los márgenes de los pétalos, células papilares en la superficie abaxial de los pétalos cerca de su base, número de haces vasculares que irrigan a cada carpelo y la existencia de una cavidad interna en la base del estilo. Los resultados permiten distinguir mejor a K. maxima de los géneros afines dentro de las Zygophyllaceae. Palabras clave: Flor; Androceo; Gineceo; Nectarios Abstract The objective is to contribute to the knowledge of the floral anatomy of Kallstroemia maxima and compare it with other related genera within the family Zygophyllaceae. Flower buds obtained from 3 plants of K. maxima were collected a day before to the anthesis. Permanent preparations were processed and observations of photomicrographs from herborizated buds were done. Anatomical descriptions of characters not previously reported are given such as: the nectaries opposed to the sepals, presence of a vascular bundle shared by a petal and the stamen opposite to it, helicoidal thickening of the epidermal cells on the petal margins, papillary cells near the base of the abaxial surface of petals, number of vascular bundles that irrigate each carpel and the existence of an internal cavity at the base of the style. The results allowed us to distinguish better Kallstroemia maxima from related genera in the Zygophyllaceae. Keywords: Flower; Androecium; Gynoecium; Nectaries ISSN versión electrónica: 2007-8706; Universidad Nacional Autónoma de México, Instituto de Biología. Open Access bajo la licencia CC BY-NC-ND (4.0) https://doi.org/10.22201/ib.20078706e.2019.90.2075 2 R.M. Fonseca et al. / Revista Mexicana de Biodiversidad 90 (2019): e902075 https://doi.org/10.22201/ib.20078706e.2019.90.2075 Introducción comparte con Kelleronia y Tribulus, entre otros caracteres, la presencia de tricomas no ramificados, vaina en los haces La familia Zygophyllaceae, considerada dentro del vasculares de las hojas, ovario sésil, ausencia de apéndices orden Zygophyllales (APG, 2016), está conformada por 22 estaminales, frutos con cocos separados en la madurez y géneros y aproximadamente 325 especies, con distribución semillas sin endospermo. tropical y subtropical asociada a zonas secas (Stevens, El estudio de la anatomía de las estructuras florales 2001). En México, se presentan 32 especies de 8 géneros es importante en la sistemática de diversos grupos de (Fagonia L., Guaicum L., Kallstroemia Scop., Larrea angiospermas (Rudall y Goldblatt, 2000; Rudall et al., Cav., Morkillia Rose et Painter, Porlieria Ruiz. et Pav., 2002). En las Zygophyllaceae se ha señalado la presencia, Sericodes A. Gray y Viscainoa Greene; Villaseñor, 2016). número, ubicación y forma de las glándulas nectaríferas Las 18 especies de Kallstroemia se distribuyen desde el como un carácter de utilidad para diferenciar taxones sur de EUA hasta Sudamérica e Indias Occidentales, 14 (Porter 1969a, 1971). Así, Chauhan (1975) consideró que de las cuales están presentes en México (Turner, 2016). la presencia de nectarios vascularizados en Balanites y Engler (1896) dividió a Zygophyllaceae en 4 no vascularizados en Tribulus contribuye a diferenciarlos. subfamilias; recientemente, con base en estudios El número de lóculos del ovario en la Zygophyllaceae moleculares se propusieron 5 grupos considerados también es un rasgo importante para caracterizar a los monofiléticos: Larreoideae, Morkilloideae, Seetzenioideae, diferentes géneros, por ejemplo Sheahan y Chase (2000) Tribuloideae y Zygophylloideae (Sheahan, 2007; Sheahan mencionaron que Augea puede distinguirse de Zygophyllum y Chase 2000). A partir de un análisis filogenético con por tener 10 lóculos; en la familia frecuentemente se secuencias moleculares (rbcL y trnL-F), Sheahan y Chase presentan de 4 o 5 lóculos, aunque pueden tener desde (2000) ubicaron a Kallstroemia dentro de la subfamilia 2 hasta 12 (Sheahan, 2007). Singh et al. (2002) también Tribuloideae y como hermano de Tribulus, Kelleronia y mencionan que la vascularización de los distintos verticilos Tribulopis, con los cuales morfológicamente comparte florales es de interés para la sistemática de esta familia. la presencia de hojas opuestas y paripinnadas con 2-12 Con base en lo expuesto anteriormente, se planteó pares de foliolos; flores pentámeras con 10 estambres realizar el estudio de la anatomía floral de Kallstroemia en 2 verticilos desiguales, ausencia de apéndices de los maxima, (L.) Hook. et Arn., para contribuir a su filamentos, frutos con 5 o 10 mericarpos indehiscentes, caracterización. semillas sin endospermo y polen poliaperturado (Sheahan, 2007). Kallstroemia se diferencia de Kelleronia porque Materiales y métodos este último género está constituido por arbustos hasta de 1.5 m, mientras que Tribulus, Tribulopis y Kallstroemia Se obtuvieron botones florales frescos y se recolectaron son hierbas postradas o ascendentes, rara vez leñosas en ramas con flores y frutos para elaborar ejemplares de su base. A su vez, Kallstroemia se distingue por presentar herbario, a partir de 3 individuos de Kallstroemia maxima, ovario con 10 lóbulos y fruto con 10 mericarpos, mientras en Yecapixtla, Morelos, México. Los ejemplares fueron que Tribulus y Tribulopis tienen ovario con 5 lóbulos y herborizados, examinados, medidos y determinados a frutos con 5 mericarpos (Sheahan, 2007). especie mediante el uso de la clave taxonómica de Porter Porter (1969a) en su tratamiento de Zygophyllaceae, (1969a). La descripción morfológica de la especie se realizó señaló que Kallstroemia, Kelleronia, Tribulopis y Tribulus midiendo y examinando los ejemplares recolectados y fue son géneros afines que se distinguen porque los estambres completada con la descripción proporcionada por Porter de la serie externa están adnados en la base a los pétalos y (1969a, b). Los ejemplares de respaldo están depositados por la presencia de tejido nectarífero entre los estambres en el Herbario de la Facultad de Ciencias (FCME) y en y el perianto. El mismo autor indicó que “entre la base de el Herbario Nacional de México (MEXU). Los botones cada filamento del verticilo interno y el sépalo opuesto a frescos fueron fijados en Navashin (Sandoval et al., 2005). éste, hay un nectario pequeño, ovoide, bilobulado”. Este material fue deshidratado en una serie de alcoholes La cercana relación entre Kallstroemia, Kelleronia, graduales en base a alcohol ter-butílico (ATB), 35%, Tribulopis y Tribulus fueron evidenciadas por diferentes 50%, 60%, 70%, 85, 95%, 100% y 3 cambios en ATB autores. Sheahan y Cutler (1993), a partir del análisis de absoluto, cada cambio con una duración de 24 horas. Se la anatomía vegetativa de 37 especies, pertenecientes a infiltró e incluyó en parafina de dureza media (punto de 19 géneros, sugirieron que, a excepción de Tribulopis, fusión 56-58 ºC). Se realizaron cortes en un micrótomo estos géneros deberían ser separados de los géneros de rotación marca American Optical modelo 820, a un zygophylloides, por lo menos, a nivel de subfamilia. grosor de 12 µm para obtener secciones transversales y Sheahan y Chase (1996) señalaron que Kallstroemia longitudinales. Las mismas fueron teñidas con safranina- R.M. Fonseca et al. / Revista Mexicana de Biodiversidad 90 (2019): e902075 3 https://doi.org/10.22201/ib.20078706e.2019.90.2075 verde rápido según Sandoval et al. (2005). Se hicieron simples, abaxialmente hirsutos, márgenes con tricomas preparaciones permanentes montadas en resina sintética. simples; nectarios 5, con forma de cojín, (fig. 1D), situados Las observaciones y fotomicrografías fueron realizadas en la cavidad que forma cada uno de los sépalos en su en un fotomicroscopio Carl Zeiss-Axioskop, con cámara base; pétalos 5, libres, convolutos, ampliamente obovados de video SONY SSC-DC544. Las imágenes se capturaron o semiorbiculares, de 4 a 9 mm de largo y hasta 7 mm de con el programa Debut y se editaron con el programa Paint ancho, amarillos o anaranjados, laminares, con una cresta Shop-Pro 8. en la cara abaxial cerca de la base, misma que se pierde Para los estudios de microscopía electrónica de hacia la parte media del pétalo; estambres 10, hipóginos, barrido
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